Middle Cambrian gogiid echinoderms from Northeast Spain: Taxonomy, palaeoecology, and palaeogeographic implications

SAMUEL ZAMORA, RODOLFO GOZALO, and ELADIO LIÑÁN

Zamora, S., Gozalo, R., and Liñán, E. 2009. Middle Cambrian gogiid echinoderms from Northeast Spain: Taxonomy, palaeo− ecology, and palaeogeographic implications. Acta Palaeontologica Polonica 54 (2): 253–265. DOI: 10.4202/app.2008.0010

Gogia parsleyi Zamora sp. nov. and Gogia sp. are described from two different echinoderm assemblages, both from the middle Cambrian of the Murero Formation (Iberian Chains, NE Spain). Gogia parsleyi is reconstructed and described on the basis of fifteen complete or partial specimens and numerous isolated plates. It is characterised by spiralled brachioles, simple epispires, sometimes covered by stereomic domes or tiny cover plates, and by thecal plates arranged in subregular circlets. This gogiid population comprises juveniles, advanced juveniles and mature individuals. The material was found in the upper part of the Murero Formation (upper Caesaraugustian–lower Languedocian). Gogia sp. is represented by two almost complete specimens and several isolated plates from the lower part of the Murero Formation (lower Caesar− augustian). The genus Gogia was first described in Western Gondwana from the Languedocian (upper middle Cambrian) of France, but the material from Spain is older and represents the oldest record of this genus in Gondwana, suggesting an early migration from Laurentia. The gogiids are well preserved in two echinoderm Lagerstätten, which, together with other echinoderms, comprise the majority of the fossil fauna. Both levels are derived from obrution deposits produced in calm and open marine conditions, sometimes affected by sporadic storms. Their holdfast morphology suggests that these gogiids were low−tier suspension feeders, living attached to trilobite fragments in a soft, muddy environment.

Key words: Echinodermata, Eocrinoidea, Gogiida, Cambrian, Murero Formation, Spain.

Samuel Zamora [[email protected]] and Eladio Liñán [[email protected]], Área y Museo de Paleontología, Departa− mento de Ciencias de la Tierra, Universidad de Zaragoza, E−50009 Zaragoza, Spain; Rodolfo Gozalo [[email protected]], Departamento de Geología. Universitat de València.C/ Dr. Moliner n° 50. E−46100 Burjasot (Valencia), Spain.

Introduction of Bohemia (Barrande 1887; Fatka and Kordule 1984, 1991), France (Ubaghs 1987) and China (Zhao et al. 1994, 2007, Echinoderms are an important component of middle Cam− 2008; Parsley and Zhao 2006). brian fossil assemblages from North Western Perigondwana. At present, only two complete and articulated eocrinoid Two problems restrict their study: (i) complete and articu− taxa are known from the Cambrian of Spain, Alanisicystis lated echinoderms are rare; and (ii) isolated plates have lim− andalusiae Ubaghs and Vizcaïno, 1991 (Fig. 1) and Ubaghsi− ited taxonomic value, usually only assignable to class level cystis segurae Gil−Cid and Domínguez, 2002. The first is from or above (Donovan and Paul 1982). Of the eleven echino− the lower Cambrian of the Ossa Morena zone (South Spain). derm classes recorded worldwide from the Cambrian (Sprin− It was originally included as a subgenus within the genus kle 1976), one of the most common is the Eocrinoidea Jaekel, Gogia, but certain features (for example the stereomic domes 1918. This Linnaean class is a paraphyletic group that com− covering complex epispires) suggest a taxonomic position prises stem−group members of the better known blastozoan independent of, but closely related to Gogia. The second, clades (Sprinkle 1973; Smith 1984; Paul 1988). Major ana− Ubaghsicystis segurae, comes from the middle Cambrian of tomical innovations of the eocrinoids include the develop− the Cantabrian zone (North Spain). It is the oldest articulated ment of erect brachioles for feeding and a long multiplated eocrinoid, with a xenomorphic stem composed of holomeric stalk, and later, a columnal−bearing stem to elevate the theca columnals (Gil−Cid and Domínguez 2002); these features sug− above the sea floor (Sprinkle and Collins 2006). gest a derived position amongst Cambrian eocrinoids. In addi− Among eocrinoids, representatives of the family Eocrini− tion to these complete eocrinoids, isolated plates belonging to dae Jaekel, 1918 are the most common, primarily from Lau− the eocrinoid Rhopalocystis? mesonesensis Clausen, 2004 rentia (Sprinkle 1973). Only a few fossils from Gondwana are from the upper lower Cambrian of the Iberian Chains, the considered as eocrinids. These are from the lower Cambrian of enigmatic blastozoan Eocystites sp. (Gil−Cid and Domínguez Spain (Ubaghs and Vizcaïno 1991), Morocco (Nardin 2006) 1998) from the middle Cambrian of Ossa Morena zone (South and China (Zhao et al. 2007), and from the middle Cambrian Spain), indeterminate eocrinoid plates (Álvaro and Vennin

Acta Palaeontol. Pol. 54 (2): 253–265, 2009 DOI: 10.4202/app.2008.0010 254 ACTA PALAEONTOLOGICA POLONICA 54 (2), 2009

for respiration, biserial brachioles and an expanded, probably short holdfast for attachment to hard substrates. The aims of this paper are twofold, firstly to describe new taxa from Spain, and secondly to analyse the geographic and stratigraphic dis− brachioles tribution of gogiids. Some preliminary data on the palaeoecol− ogy of eocrinoids and other echinoderms from the Murero Formation is also discussed. Institutional abbreviation.—MGM, Museo Geominero, Ma− drid, Spain; MPZ, Museo de Paleontología, University of Zaragoza, Spain.

Geological setting and stratigraphy

The Murero Formation is exposed in several Cambrian local− ities (Fig. 2) in the Iberian Chains (NE Spain), all of which are rich in trilobites, echinoderms, brachiopods, and Burgess Shale−type fossils. Trace fossils are also present. Two large anal pyramid Palaeozoic outcrops trending NW−SE, separated by the Ter− tiary Calatayud−Teruel basin, constitute the central part of the Iberian Chains. In this region, Palaeozoic rocks are struc− tured into three tectonostratigraphic units, the Mesones Unit, Herrera Unit and Badules Unit (Gozalo and Liñán 1988), and theca Cambrian rocks are common in all three. The Cambrian stratigraphy of the Iberian Chains was es− tablished by Lotze (1929); the lithostratigraphic nomencla− ture has subsequently undergone minor modifications (Lotze 1958, 1961; Schmitz 1971; Liñán et al. 1992; Álvaro 1995). The units in this sequence (in ascending stratigraphic order) are: the Bámbola Formation, Embid Formation, Jalón For− mation, Ribota Formation, Huérmeda Formation and Daroca 5mm Formation for the lower Cambrian; the Mesones Group (Val− holdfast demiedes, Mansilla and Murero formations) for the lower– middle Cambrian; and the Acón Group for the middle Cam− Fig. 1. Holotype (MGM2005K) of gogiid echinoderm Alanisicystis andalu− brian–Furongian. The palaeontological contents of each unit siae Ubaghs and Vizcaïno, 1991 from the lower Cambrian of the Ossa were summarised by Liñán et al. (1996, 2002) and Gozalo et Morena zone (South Spain). Photograph of latex cast whitened with NH4Cl. al. (2008). The fossils described herein came from two different local− 1997) from the middle Cambrian (Mansilla Formation, Ibe− ities, Murero and Purujosa villages; both are situated in the rian Chains) and columnals of uncertain affinity from the Badules unit, which is considered to be the prolongation of the Furongian (Acón Group, Iberian Chains) (Zamora et al. in West Asturian−Leonese Zone to the southeast (Fig. 2A). They press), have also been described. are from two different beds in the Murero Formation. This for− Echinoderms from the Murero Formation are extremely mation represents a monofacial deposit of green lutites with diverse, with a long and complete fossil record (Zamora et al. interbedded carbonate nodules and very fine sandstones. The 2007), but few have been formally described so far (Friedrich original sediments were deposited in a shallow marine envi− 1993; Zamora and Rahman 2008). In this paper we report two ronment (sublittoral facies, sensu Liñán 1995). The Dyctio− new gogiid eocrinoids from the middle Cambrian Murero For− nina–Acrothele brachiopod assemblage is well represented mation. Gogia parsleyi Zamora sp. nov. and Gogia sp. are the throughout the formation and indicates the predominance of first representatives of the group reported from the Iberian low−energy deposits (Liñán and Mergl 2001). Chains. Gogia sp. is also the oldest representative of this genus The presence of agnostoid and polimeroid cosmopolitan in Western Gondwana and suggests taxa migrated to Gond− trilobites suggests an outer sublittoral facies for the Murero wana from the Laurentian palaeocontinent. Formation. These eocrinoids possess some plesiomorphic (primitive) Locality 1.—Murero is a small village situated on the west− characters, including an irregularly plated theca with larger ern branch of the Iberian Chains, located 80 km south of primary and smaller secondary plates, epispires (sutural pores) Zaragoza (Fig. 2B). The Cambrian rocks in this area show a ZAMORA, GOZALO, AND LIÑÁN—CAMBRIAN GOGIID ECHINODERMS FROM SPAIN 255

Moncayo Jarque Fault Herrera Unit Badules Unit Mesones Unit Precambrian rocks

Tablado Range Tabuenca Borobia Purujosa Zaragoza

Mesones de Isuela

La Almunia de Da Godina

Calatayud -Teruel Basin

Calatayud Datos Fault Puig Moreno

Herrera de los Villafeliche Navarros

oo Murero Long. W 005 Long. 000 Badules C OC Daroca WAL

GC

Lat. N 41o

Iberian Chains ELA

OM Calamocha 200 km SP

010km Pre-Hercynian Montalbán Lat. N 36o outcrops

Fig. 2. Geological setting of the two discussed localities in the Iberian Chains (after Liñán et al. 2008). A. Pre−Hercynian outcrops and tectono−stratigraphic zones of the Iberian Peninsula; the Iberian Chains are framed. Zones: CZ, Cantabrian; WALZ, West Asturian−Leonese; GCZ, Galician−Castilian; ELAZ, East Lusitanian−Alcudian; OMZ, Ossa−Morena Zone; SPZ, South Portuguese. B. Pre−Hercynian outcrops and tectono−stratigraphic zones and units of the Iberian Chains; Murero and Purujosa (indicated by stars) (Modified from Gozalo and Liñán 1988). lower and middle Cambrian sequence divided into Valde− nus (Sdzuy, 1968), Badulesia granieri (Thoral, 1935), Cono− miedes, Mansilla, and Murero formations in a normal suc− coryphe (Parabailiella) languedocensis Thoral, 1946 and cession dipping to the south (Liñán and Gozalo 1986). Eo− Condylopyge sp. The presence of both agnostoid and poli− crinoids were collected in the upper part of level 12 of meroid trilobites suggests open marine conditions; however, Rambla de Valdemiedes 1 section, which represents the base the sponge Leptomitus conicus is also present, which is typi− of the Murero Formation (Fig. 3). This level is five metres cal of soft substrates in low or moderately low energy condi− thick, comprising green shales and carbonate nodules (see tions (García−Bellido et al. 2007). Liñán and Gozalo 1986, and García−Bellido et al. 2007), and containing trilobites, echinoderms, brachiopods, sponges, Locality 2.—Purujosa is located in the Moncayo Natural and ichnofossils. The presence of the trilobite Badulesia Park at the Tablado Range, 75 km west of Zaragoza in the granieri (Thoral, 1935), indicates that the level belongs to north of the Iberian Chains (Fig. 2B). Eocrinoids were col− the Badulesia granieri Zone (lower Caesaraugustian). lected from the top of the Murero Formation in the Puru− The eocrinoids from this locality were found in a thin josa−4 section (Fig. 3); in the study area, this formation oc− level associated with the trilobites Eccaparadoxides asturia− curs as a 75 m sequence of shales, nodular carbonates and

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Fm. RV 1Stage Pur 4 Fm. Barrande, 1860), Peronopsis acadica (Hart, 1868), Perono− psis ferox (Tullberg, 1880), Solenopleuropsis simula Sdzuy,

BOROBIA 1958, S. marginata Sdzuy, 1958, S. thorali Sdzuy, 1958,

BOROBIA Conocoryphe (Conocoryphe) heberti Munier−Chalmas and Bergeron, 1889, and Ctenocephalus aff. coronatus (Bar− rande, 1846); these taxa suggest a lower Languedocian age (middle Cambrian). Thus, the age of the echinoderm level Languedocian containing eocrinoidean fossils is likely to be uppermost Caesaraugustian or lowermost Languedocian. The oldest cinctan echinoderm is also reported from this section, in the underlying Mansilla Formation (Rahman and Zamora in press).

Systematic palaeontology (by Samuel Zamora)

MURERO MURERO Phylum Echinodermata Bruguière, 1791 (ex Klein, upper 1734) Caesaraugustian Subphylum Blastozoa Sprinkle, 1973 Class Eocrinoidea Jaekel, 1918 Order Gogiida Broadhead, 1982 Family Eocrinidae Jaekel, 1918 Genus Gogia Walcott, 1917 Type species: Gogia prolifica Walcott, 1917, lower Middle Cambrian, middle British Columbia (Canada). Caesaraugustian Discussion.—Gogia is the most abundant and diverse Cam− brian eocrinoid (Sprinkle 1973). Thirteen species have been described in the lower and middle Cambrian of Laurentia; lower only one of these, Gogia ojenai Durham, 1973, is from the Caesaraugustian lower Cambrian (Robison 1965; Sprinkle 1973; Sprinkle and MANSILLA Collins 2006). There are, however, several new species from 20 m the lower Cambrian of Laurentia that still require formal de− scription (Bryan C. Wilbur, personal communication 2005).

MANSILLA Until now, the first convincing appearance of Gogia in West− ern Perigondwana was in the upper middle Cambrian of Leonian 10 Montagne Noire (France), where Gogia gondi Ubaghs, 1987 is reported. The Spanish material described here suggests an 0 older first occurrence of the genus Gogia in Gondwana, in carbonate nodules dolomarly shale sandstone the lower middle Cambrian. One point of discussion within the genus Gogia is the sub− dolostone shale fault genus Alanisicystis Ubaghs and Vizcaïno, 1991, from the Fig. 3. Sections of the middle Cambrian Murero Formation in Murero and lower Cambrian of Southern Spain (Fig. 1). This subgenus is Purujosa indicating the levels with Gogia sp. and Gogia parsleyi Zamora characterised by “single or partioned epispires provided with sp. nov. external dome−like stereomic cover”. There are no other Gogia species that show this peculiar type of epispire (com− fine sandstones. Fossils were collected in a thin (35 cm) level plex and covered) (Sprinkle 1976; Ubaghs 1987). For this rea− of green−grey shale, rich in cinctans (Gyrocystis platessa son, we believe that Alanisicystis Ubaghs and Vizcaïno, 1991 Jaekel, 1918) and eocrinoideans. Trilobite fragments and should be a separate genus rather than a subgenus of Gogia trace fossils are also present, but so far only the trilobite (note, type material of Alanisicystis andalusiae Ubaghs and Eccaparadoxides brachyrachys (Linnarsson, 1883) has been Vizcaïno, 1991 previously deposited in Carcassone, France identified. The trilobite assemblage in the overlying level is [Vizcaïno collection numbers: VCE 11, 1–3, 23, 24, 25, 26] is composed of Eccaparadoxides? pradoanus (Verneuil and now deposited in the MGM. New collection numbers are ZAMORA, GOZALO, AND LIÑÁN—CAMBRIAN GOGIID ECHINODERMS FROM SPAIN 257

5mm

2mm 5mm

2mm 1mm

5mm 5mm

5mm 2mm

2mm 2mm

Fig. 4. A–E. Eocrinoid blastozoan Gogia parsleyi Zamora sp. nov. A. Paratype MPZ2006/556a, b; part (A1) and counterpart (A2) of a slightly disturbed small specimen attached to a free cheek of Eccaparadoxides sp. fragment. The arrow indicates where the holdfast attaches to the trilobite element. B. MPZ2006/559b; accumulation of disarticulated plates from eocrinoids (Gogia parsleyi Zamora sp. nov.) and cinctans. C. Paratype MPZ2006/557a, b; nearly complete specimen with thecal plates slightly disturbed. D. Paratype MPZ2004/215; partial theca with ornamented plates and a very short holdfast (indicated by the arrow). E. Paratype MPZ2004/214; specimen with an almost complete theca. F, G. Gogia sp. F. MPZ2004/194a; partly complete speci− men with a possible periproct on the lateral surface (see white arrow). G. MPZ2004/195a, b; partially disarticulated specimen (G1), counterpart of the same specimen (G2), detail of two adjoined plates with epispires (G3). Photographs of latex casts whitened with NH4Cl.

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5mm

Fig. 5. Slab with two nearly complete, articulated and exquisitely preserved specimens of eocrinoid blastozoan Gogia parsleyi Zamora sp. nov., with some isolated plates belonging to cinctan carpoids. Both specimens probably represent an early mature stage (TH = 12 mm). The left specimen (holotype MPZ2004/162a) shows many of the diagnostic features referred to in the text. The right specimen (paratype MPZ2004/161a) shows the holdfast separated from the theca (indicated by an arrow). Photograph of latex cast whitened with NH4Cl. MGM2003K 1–3, MGM2004K, MGM2005K, MGM2006K, Almost no stalk, except for an expanded holdfast composed of MGM2007K). tiny globular plates. Material.—14 articulated and nearly complete specimens (ten Gogia parsleyi Zamora sp. nov. with both part and counterpart), MPZ2004/161–163, 214, Figs. 4A–F, 5–9. 215, 238; MPZ2006/556–558, 560; MPZ2008/160–162, 164. Etymology: In honour of Prof. Ronald L. Parsley (Tulane University, Two disarticulated specimens, MPZ2006/559 and MPZ2008/ New Orleans), for his contributions to the understanding of gogiid 163, and 23 isolated plates (MPZ2004/216–237, 239). All fos− eocrinoids and Palaeozoic echinoderms in general. sils are preserved as natural moulds in a grey−green shale. Ar− Type material: Holotype: MPZ2004/162a, b, almost complete specimen ticulated specimens with delicate, intact structures suggest without holdfast (Fig. 5). Paratypes: MPZ2004/161, 163, 214, 215, 238; rapid burial while alive by a storm−induced obrution deposit. MPZ2006/556–558, 560; MPZ2008/160–164, complete or partial spec− imens. Description.—The shape of the theca is ellipsoidal to roun− Type locality: La Borraca Creek, four kilometers to the southwest of ded; the size of the holotype theca is 12 × 11 mm (Fig. 5). All Purujosa village (Zaragoza, NE Spain) in the Moncayo Natural Park. the recovered specimens have been flattened by collapse Type horizon: Murero Formation, uppermost Caesaraugustian or lower− and/or compaction. There are 35 thecal plates per exposed most Languedocian, Middle Cambrian. side, which exhibit polygonal outlines and subregular ar− Diagnosis.—Eocrinid with globular−shaped theca and large, rangements in 6 or 7 circlets; a number of the plates are large slightly convex thecal plates (slightly ornamented). Epispires (up to 3 mm) (see reconstruction, Fig. 6). There is no clear are simple and numerous, with a well developed raised rim gradient in the size of the plates, but the larger ones are con− sometimes covered by a stereomic dome or by small plates. centrated in the lower–middle portions of the theca; plates Thecal plates are arranged into six or seven subregular circlets. decrease in size aborally and especially adorally. Additional At least seven long, narrow biserial and spiralled brachioles. plates are intercalated along sutures, as shown by the juvenile ZAMORA, GOZALO, AND LIÑÁN—CAMBRIAN GOGIID ECHINODERMS FROM SPAIN 259 specimen MPZ2006/556 (Fig. 4A). Plates are slightly domed on their external surface and are typically ornamented with tiny granules (Fig. 4D). On the internal surface, they are unornamented and slightly concave (Fig. 4B). Some stereo− mic structures are well preserved (Fig. 7). Sutural pores (epispires) occur over the entire theca, with a regular distribution and size (Figs. 5, 7B, C, 8A, B, 9C). Epispires are surrounded by a prominent raised rim (Figs. 6, 7C, 8E, F, 9C), which usually crosses the associated plate su− ture(s). Epispires are present on the sutures between two plates or at the corners between three plates (Fig. 8F); and they are sometimes more developed in one of the two adjacent plates. The pores show an almost uniform diameter. The number of epispires per plate ranges from 4 (MPZ2004/162a) to 17 (MPZ2004/217) and there are up to four epispires per plate side. Some epispires show an external dome−like stereomic cover (Figs. 6, 8E), others are covered by numerous tiny plates (Figs. 4E, 8F). The covered epispires are only present in the lower–middle theca and sometimes these structures do not cover the whole pore. These epispires are rounded to elliptical and have an epispire H/W ratio varying from 1 to 3. Brachioles are attached in groups of two to a modified single thecal plate, which is projected at the periphery of the oral region (Fig. 8C). They are spiralled with a quite loose twist (approximately 0.5 mm) (Fig. 8G) and are sometimes coiled at their tips (Figs. 5, 8D). The length of the brachioles is at least 25 mm, twice the length of the theca. Seven brachioles are preserved in the most complete specimens (MPZ2004/162a), there were probably more in living ani− Fig. 6. Reconstruction of eocrinoidean blastozoan Gogia parsleyi Zamora mals. Brachioles are biserial, with the alternating brachiolar sp. nov. (by Santiago Alberto, after a sketch by SZ), based on the holotype plates numbering about three or four per millimetre (Figs. MPZ2004/162a. The holdfast, not preserved in the holotype, is recon− 8A, G, 9A). Cover plates are not well preserved; there are structed based on paratypes MPZ2004/161 and MPZ2004/215.

1mm

Fig. 7. Details of element structures in eocrinoidean blastozoan Gogia parsleyi Zamora sp. nov. (SEM photos of latex casts). A. Fragment of a holdfast from the specimen MPZ2004/161, consisting of an aggregate of globular plates (A1), details of specimen (A2,A3). B. Internal view of a plate showing sutural pores of epispires (MPZ2004/225); general view of an isolated plate (B1), detail of the epispire (B2). C. External surface of a plate (MPZ2004/232) (C1), details of the epispires showing the raised rim and stereomic structures (C2, C3). Arrows indicate enlarged details.

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1mm

1mm 1mm

1mm

3mm

Fig. 8. Camera lucida drawings of eocrinoid blastozoan Gogia parsleyi Zamora sp. nov. A. General view of the paratype MPZ2004/161, detached holdfast below. B. Thecal plate, half−epispires with their prominent rim indicated. C. Two brachioles on a single thecal plate. D. Biserial brachiole terminally en− rolled. E. Epispire covered by a single domal plate. F. Epispire covered by tiny plates. G. Spiralled brachiole. two on a brachiolar plate, with a spinous projection in lateral kle, personal communication 2008). Gogia parsleyi Zamora view. The preservation of the brachioles in the holotype sug− sp. nov. is similar to Gogia hobbsi Sprinkle, 1973, particularly gests that they were flexible. in the ornamentation of the thecal plates and the spiralled The holdfast is preserved in some specimens, but no stalk brachioles; however, Gogia hobbsi has a conical thecal shape, is apparently present. In MPZ2004/161a (Fig. 5) the holdfast is smaller in size with fewer plates and has a stalk. appears disarticulated, in specimens MPZ2004/215 (Fig. Gogia parsleyi is likely to be closely related to Alanisi− 4D) and MPZ2008/162 (Fig. 9A) it is apparently still con− cystis Ubaghs and Vizcaïno, 1991, because both have stereo− nected to the base of the theca. It is composed of numerous mic cover domes on some epispires in the lower part of the tiny, globular plates, which are clearly visible in SEM im− theca, but it differs from this species by having simple epi− ages (Fig. 7A). The transition between the holdfast and theca spires and plate ornamentation composed of a slightly rough is abrupt, consisting of a change in the type and size of plates. surface texture. Moreover, in Gogia parsleyi some epispires are covered by tiny plates rather than domes, which are Discussion.—Gogia parsleyi Zamora sp. nov. conforms to always observed in Alanisicystis (Fig. 1). This feature is the diagnosis proposed by Walcott 1917 (see also, Sprinkle equally well shown in Rhopalocystis destombesi Ubaghs, 1973; Sprinkle and Collins 2006) for the genus Gogia. It has 1963, in which a group of small plates covers the sutural a unique character combination of simple covered epispires; pores (Ubaghs 1963: fig. 8.5–7) and in Globoeocrinus glo− large thecal plates arranged in subregular circlets; spiralled bulus Zhao, Parsley, and Peng, 2008. A new gogiid from brachioles; and the absence of a stalk connecting the ex− Morocco that is under description (Nardin 2006) shares panded holdfast with the theca. many features with Gogia parsleyi, such as the presence of The general body structure of Gogia parsleyi Zamora sp. covered epispires, but they differ mainly in the type of nov. (Fig. 6) shows many features that are shared with other epispires, which are simple in the Spanish material and com− gogiids, such as erect biserial brachioles, thecal plates ar− plex in the Moroccan specimen (Elise Nardin, personal com− ranged in poorly developed circlets, epispires and an attach− munication 2006). ment appendage consisting of only a holdfast. Sinoeocrinus Zhao, Huang, and Gong, 2004 differs by This species differs from other Cambrian gogiids in sev− lacking the stereomic domes or tiny plates covering epispires. eral characteristics; for example, thecal plates organised into Gogia parsleyi differs from Akadocrinus Prokop, 1962 in the subregular arranged circlets and the presence of covered epi− stalk, which is nearly absent in the former and composed of spires. One undescribed species of Gogia from North America many fusular rings in the later. Furthermore, they differ in gen− also shows stereomic domes covering epispires (James Sprin− eral thecal shape. ZAMORA, GOZALO, AND LIÑÁN—CAMBRIAN GOGIID ECHINODERMS FROM SPAIN 261

A B C

3mm D

3mm 4mm 4mm

Fig. 9. Eocrinoid blastozoan Gogia parsleyi Zamora sp. nov. A. Paratype MPZ2008/162. Complete juvenile specimen (length of theca is about 5 mm); brachioles are spiralled and probably show a 2−2 pattern. B. Paratype MPZ2006/558a. Advanced juvenile specimen (length of theca is 11 mm ). C. Paratype MPZ2008/164b. Fragment of theca with tessellated plates, epispires with characteristic rim. D. Paratype MPZ2008/161, upper part of theca. Photographs are of latex casts taken from natural moulds whitened with NH4Cl.

Marjumicystis Ubaghs and Robison, 1985 also appears to Stratigraphic and geographic distribution.—Upper part of lack a stalk, but differs principally in lacking well developed Murero Formation, uppermost Caesaraugustian or lower− epispires. most Languedocian (middle Cambrian). Parsley and Zhao (2006) provided a complete ontogenetic sequence for the Cambrian eocrinoid Sinoeocrinus lui Zhao, Gogia sp. Huang, and Gong, 2004, describing, for the first time, the Fig. 4F, G. ontogenetic changes occurring in a gogiid population. These ontogenetic stages were described in terms of thecal height Material.—Two incomplete specimens partially articulated (TH) and were thus divided into juvenile, advanced juvenile, (MPZ2004/194, 195) and several isolated plates that probably mature and advanced mature, or gerontic, stages. The material belong to a single disarticulated specimen (MPZ2004/196). of Gogia parsleyi described in this paper is not very abundant, All the material is preserved as natural moulds coated with li− but a few different ontogenetic stages can be recognised; these monite in green shale. stages are important to understand feature changes in speci− Description.—The most complete specimen (Fig. 4F) mens of different sizes. The TH of specimens ranges from 5 (MPZ2004/194) possesses a rounded, conical theca, with a mm to 12 mm. The smallest specimens (Figs. 4A, 9A), with a narrow base and expanded adoral surface. The theca height/ TH of 6 mm and 5 mm, respectively, probably represent juve− width ratio is approximately 2; the number of plates is ap− niles or early advanced juveniles (sensu Parsley and Zhao proximately 70 per exposed surface. Plates are irregular in 2006). They have a 2−2 ambulacral pattern (Fig. 9A), lack of shape and ornamented with small crests; they are irregu− ornamentation and very small epispires without a raised rim. larly arranged with the primary plates surrounded by many The specimen which is intermediate in size, with a TH of ca. small secondary ones. Plates are thick compared to those of 7.5 mm (Fig. 9B), shows intermediate features, lacking orna− other gogiids. Epispires are rare and confined to larger mentation and possessing an incipient raised rim bordering plates. They are small and lack a raised rim (Fig. 4G3), with epispires, which are very small. It probably represents an ad− a height/width ratio of about 1.7. A rounded lateral opening vanced juvenile. The largest specimens have a TH of ca. 12 is observed in one specimen, with a diameter of 0.65 mm mm and show some very distinctive features. They possess (Fig. 4F), which corresponds (in position) to the anal pyra− more than five brachioles, probably in a 2−1−2 pattern, orna− mids of other gogiids. mented thecal plates and well developed, sometimes covered Brachioles are disarticulated in two of the specimens; as a epispires with prominent raised rims. All these features are result it is impossible to know their exact number and de− probably indicative of early mature specimens. tailed morphology. They were probably quite low in number

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Mediterranean Subdivision Czech South China USA and Canada Republik Area proposal of ISC

Ellipsocephalus hoffi– Rejkocephalus Zone Languedocian Paradoxides gracilis Wangcunian taxon range Zone Marjuman Drumian Eccaparadoxides pusillus–Paradoxides gracilis interval Zone O. hybridus Caesar- taxon range Zone Augustain Eccaparadoxides pusillus taxon range Zone Taijiangian Topazan Stage 5 Ornamentaspis– Conocoryphe horizon Leonian

Delmaran

Bilbilian

Duyunian Stage 4 Dyerian Marianian

Nangaoan (p.)

Fig. 10. Stratigraphic distribution of gogiids (indicated by stars) in the early and middle Cambrian. The biostratigraphy for the Czech Republic is based on Fatka (2006); the stages for the Mediterranean area are from Liñán et al. (2002) and Gozalo et al. (2008); the stages for South China are taken from Peng and Babcock (2001) and Peng (2003); the stages for USA and Canada (Laurentia) are from Sundberg and McCollum (2003) and Sundberg (2005). The Subdivi− sion proposal of ISC for the Cambrian System is from Zhu et al. (2006).

(4?), and fairly long and large. The brachiolar plates show a occur in the USA; there is a nearly continuous sequence of well developed food groove. different Gogia species from the basal Delamaran to the mid− Discussion.—The general morphology of the theca, disposi− dle Marjuman (stages from the North American chrono− tion of plates and the structure of epispires suggest that the stratigraphic scale, see Sprinkle 1976). This extensive record material should be included in the genus Gogia. strongly suggests that North America was the area where Stratigraphic and geographic distribution.—Lower part of Gogia first evolved, although it probably migrated, at least the Murero Formation, Badulesia granieri Zone, lower once, to northern Perigondwana (France, Ubaghs 1987; and Caesaraugustian. Spain, herein). All other gogiids found in parts of Gondwana are middle Cambrian (Fig. 10), these generally occur as isolated records Palaeobiogeography and of single species in different levels. The earliest are Sinoeo− crinus lui Zhao, Huang, and Gong, 1994 and Globoeocrinus palaeoecology of gogiids globulus Zhao, Parsley, and Peng, 2008, from the base of the Taijiangan stage of China (Peng and Babckock 2001; Peng Stratigraphic occurrence and palaeobiogeography.—The 2003). The next representative, in ascending stratigraphical oldest known gogiids were reported from the early Cambrian order, is Gogia sp., from the lower Caesaraugustian of Spain of California, USA (see Sprinkle 1976), Ossa Morena, Spain (herein). Finally, the latest occurrences are known from four (Ubaghs and Vizcaïno 1991), Anti−Atlas, Morocco (Nardin horizons in the Czech Republic (Fatka 2006), France (Ubaghs 2006), and China (Zhao et al. 2007); all of these are approxi− 1987), and Spain (herein); these horizons all have a similar mately the same age (Fig. 10). The next oldest records also age, uppermost Caesaraugustian to lowermost Languedocian ZAMORA, GOZALO, AND LIÑÁN—CAMBRIAN GOGIID ECHINODERMS FROM SPAIN 263

NORTH LAURENTIA CHINA

SOUTH CHINA SIBERIA

GONDWANA

BALTICA AVALONIA Lower Cambrian Gogiids Middle Cambrian Gogiids

Fig. 11. Palaeogeographic distribution of gogiids in the early and middle Cambrian; reconstruction after McKerrow et al. (1992).

(Fig. 10). The youngest gogiids from Europe and North Amer− isolated echinoderm plates appear unabraded, perhaps indi− ica are similar in age. cating a time averaging process, compaction of the surround− Gogiids are known from three distinct biogeographic ar− ing sediment or the action of infaunal animals. Further work eas: the Rocky Mountains (western North America), with the is needed to explain the association of both articulated speci− exclusive presence of the genus Gogia; northern Perigod− mens and disarticulated plates. Similar modes of origin have wana (Europe and North Africa), where Gogia and another been proposed for other Cambrian echinoderm assemblages three genera have been found; and South China, where three (Sprinkle 1976; Bell and Sprinkle 1978; Ubaghs and Robi− additional genera are reported. son 1985; Ubaghs 1987; Friedrich 1993; Lin et al 2008). The distribution of gogiids (Fig. 11) is apparently re− Locality 1, in the lower part of the Murero Formation stricted to tropical and subtropical regions when we plot their (Badulesia granieri Zone), is a clay−shale containing a fossil occurrences on the Cambrian palaeogeographic reconstruc− assemblage made up of abundant echinoderms, sponges and tion of McKerrow et al. (1992). This pattern has also been de− trilobites. The echinoderms include the eocrinoid Gogia sp., scribed for other groups, including trilobites, bradoriids, and the enigmatic blastozoan Eocystites sp., and a new cinctan demosponges, and could be interpreted as a climatic control carpoid. on the geographic distribution of gogiids. Locality 2 is located in the upper part of the Murero For− Another suggestion is that two main lineages existed; one mation (uppermost Caesaraugustian or lowermost Languedo− is the genus Gogia that evolved in the Rocky Mountains, cian). The echinoderms are the eocrinoid Gogia parsleyi, the which migrated to Europe during the middle Cambrian. The cinctan Gyrocystis platessa Jaekel, 1918 and isolated plates other lineage comprises genera of gogiids that are exclu− that probably belong to the stylophoran carpoid Ceratocystis? sively from Gondwana regions. As has been suggested for other Cambrian echinoderm groups (Lefebvre 2007), dense assemblages of echinoderms Palaeoecology.—The gogiids from the Iberian Chains pro− seem relatively incompatible with a rich and diverse associ− vide a good opportunity to document the palaeoecology of ated fauna. This is illustrated in the two assemblages de− this group in the Perigondwanan margin during the middle scribed herein, where echinoderms are the major fossil group Cambrian. Fossil echinoderms from the two studied beds are and trilobites show little diversity. Furthermore, the com− exquisitely preserved, with some examples of complete and plete absence of phosphatic brachiopods and the scarcity of articulated eocrinoids with the brachioles still attached, and agnostoid trilobites, which are common in other levels of the the theca and holdfast intact; furthermore, cinctan carpoids Murero Formation, may imply relatively shallow water con− are preserved with the delicate labrum in place and with com− ditions for the two echinoderm assemblages. plete steles. All these features of excellent preservation are Different strategies of attachment have been proposed for associated with the same lithology type, suggesting that primitive eocrinoids living on this poorly consolidated sedi− echinoderms were living together and killed, probably by ment (Sprinkle 1973; Parsley and Prokop 2004; Sprinkle and obrution processes caused by storms. In all cases, abundant Wilbur 2005; Dornbos 2006). The most common hypothesis

http://app.pan.pl/acta54/app54−253.pdf 264 ACTA PALAEONTOLOGICA POLONICA 54 (2), 2009 is attachment by bioglue to hard elements, such as brachio− Texas, Austin, USA) and Reimund Haude (University of Göttingen, pods or trilobite carapace fragments. Gogiids from the mid− Germany), made many useful suggestions for improvement. The re− dle Cambrian of China recently provided further information search was partly supported by the Project BTE2006−12975 of the Spanish Ministerio de Ciencia y Tecnología, and by the Grupo Conso− on attachment strategies. Based on a large collection, Lin et lidado E−17 and PM067/2006 of the Aragón Government. SZ holds a al. (2008) indicated that 73% of gogiids are preserved at− pre−doctoral research grant from the Departamento de Educación, tached to skeletal substrates. One specimen of G. parsleyi ap− Cultura y Deporte of the Aragón Government. pears to be fixed to a trilobite fragment (Fig. 4A), in this case a librigena of Eccaparadoxides sp.; the attachment structure of this specimen is composed of numerous tiny globular plates that make a holdfast. References In summary, the eocrinoids described herein were one of the most important components of the soft−bottom communi− Álvaro, J.J. 1995. Propuesta de una nueva unidad litoestratigráfica para el Cámbrico Medio−Superior de las Cadenas Ibéricas (NE España): El ties that lived in the calm, shallow, open marine conditions Grupo Acón. Boletín de la Real Sociedad Española de Historia Natural associated with the Murero Formation. They coexisted with (Geología) 90: 95–106. other echinoderms that make up the most important compo− Álvaro, J.J. and Vennin, E. 1997. Episodic Development of Cambrian nent of these fossil assemblages within eocrinoids. Other fos− Eocrinoid−Sponge Meadows in the Iberian Chains (NE Spain). Facies sils associated with echinoderms include trilobites, sponges, 37: 49–64. Barrande, J. 1887. Système Silurien du centre de la Bohême. Vol. VII. Classe and small shelly fossils, which are always a minor part of the des échinodermes, ordre des Cystidées. 233 pp. Rivnac−Gerhard, Prague. community. This peculiarity has been observed in other Bell, B.M. and Sprinkle, J. 1978. Totiglobus, an unusual new edrioasteroid Cambrian and lower Ordovician areas where echinoderms from the Middle Cambrian of Nevada. Journal of Paleontology 52: are the most important fossil group (Sprinkle 1973; Sprinkle 243–266. 1976; Ubaghs and Robison 1985; Lefebvre 2007). The most Donovan, S.K. and Paul, C.R.C. 1982. Lower Cambrian echinoderm plates from Comley, Shropshire, England. Geological Magazine 119: 611–614. common feature of the two echinoderm assemblages de− Dornbos, S.Q. 2006. Evolutionary palaeoecology of early epifaunal echino− scribed herein is that eocrinoids commonly show low diver− derms: Response to increasing bioturbation levels during the Cambrian sity, being represented by one taxon per bed. When other radiation. Palaeogeography, Palaeoclimatology, Palaeoecology 237: echinoderms appear, they show a distinct body plan that sug− 225–239. gests a different ecological niche. “One−species−to−one−lo− Fatka, O. 2006. Biostratigraphy of the Jince Formation (Middle Cambrian) in the Príbram−Jince Basin: Historical review. Acta Universitatis Carolinae cality” (sensu Sprinkle 1976) is the most common situation – Geologica 47: 53–61. among eocrinoids from the early–middle Cambrian of Lau− Fatka, O. and Kordule, V. 1984. Acanthocystites Barrande, 1887 (Eocri− rentia (Sprinkle 1976; Ubaghs and Robison 1985), but the noidea) from the Jince Formation (Middle Cambrian) of the Barrandian data reported in this paper (as well as previous works by area. Věstník Ústředního ústavu geologického 59: 299–302. Ubaghs [1987] and Ubaghs and Vizcaïno [1991]) confirms Fatka, O. and Kordule, V. 1991. Akadocrinus knizeki sp. nov., gogiid this scenario also for Gondwana. Perhaps this comes from eocrinoid from Czechoslovakia (Echinodermata, Middle Cambrian). Věstník Ústředního ústavu geologického 66: 239–243. the apparent gregarious nature of the individuals in each spe− Friedrich, W.P. 1993. Systematik und Funktionsmorphologie mittelkambri− cies, which exclude other species from a single “garden” scher Cincta (Carpoidea, Echinodermata). Beringeria 7: 1–190. or colony (Sprinkle 1976). Finally, the association of differ− García−Bellido, D.C., Gozalo, R., Chirivella Martorell, J.B., and Liñán, E. ent types of echinoderms in the same levels was probably 2007. The Demosponge genus Leptomitus and a new species from the controlled for the relatively similar ecological requirements Middle Cambrian of Spain. Palaeontology 50: 467–478. Gil Cid, M.D. and Domínguez, P. 1998. “Carpoidea” and Pelmatozoa from (Lefebvre 2007). the Middle Cambrian of Zafra (SW Spain). In: R. Mooi and M. Telford (eds.), Echinoderms: San Francisco, 93–98. A.A. Balkema, Rotterdam. Gil Cid, M.D. and Domínguez, P. 2002. Ubaghsicystis segurae nov. gen. y sp., nuevo Eocrinoide (Echinodermata) del Cámbrico Medio del Norte Acknowledgments de España. Coloquios de Paleontología 53: 21–32. Gozalo, R. and Liñán, E. 1988. Los materiales hercínicos de la Cordillera We thank Ronald L. Parsley (Tulane University, New Orleans, USA) Ibérica en el contexto del Macizo Ibérico. Estudios geológicos 44: 399–404. not only for accepting the dedication of the new species described Gozalo, R., Liñán, E., Gámez Vintaned, J.A., Dies Álvarez, M.E., Chirivella herein, but also for reviewing this paper and making useful comments. Martorell, J.B., Zamora, S., Esteve, J., and Mayoral, E. 2008. The Cam− We also thank Elise Nardin (University of Rennes, France) for sending brian of the Cadenas Ibéricas (NE Spain) and its trilobites. Cuadernos latex casts of French specimens and comments on Moroccan eocri− del Museo Geominero 9: 137–151. noids, Daniel Vizcaïno (Carcassone, France) for his comments on the Lin, J.−P., Ausich, W.I., Zhao, Y.L., and Peng, J. 2008. Taphonomy, palaeo− Cambrian stratigraphy of Montagne Noire (France) and Bryan C. Wil− ecological implications, and colouration of Cambrian gogiid echinoderms bur (University of Texas, Austin, USA) for interesting discussions from Guizhou Province, China. Geological Magazine 145: 17–36. about lower Cambrian eocrinoids from the western USA. We thank Lefebvre, B. 2007. Early Palaeozoic palaeobiogeography and palaeoecol− Isabel Pérez (Zaragoza University, Spain) for photographs and help us ogy of stylophoran echinoderms. Palaeogeography, Palaeoclimato− with Fig. 3. 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